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Armand De Ricqles Biologie historique et Évolutionnisme M. Armand DE RICQLÈS, professeur ENSEIGNEMENTS 1. Au titre du Collège de France 1.a. — Cours (les vendredis 10, 17, 24, 31 janvier, 7, 14, 21, 28 février, 7 et 14 mars 2003, soit 20 heures) Crocodiles, Dinosaures, Oiseaux : le « Parc Jurassique » revisité (suite). Poursuivant cette année l’analyse de l’évolution et de la paléobiologie des Archosauriens, je me suis proposé de présenter un panorama aussi complet et précis que possible des Sauropodomorphes, le groupe-frère des dinosaures bipèdes carnivores ou Théropodes passés en revue l’année dernière. Les Sauropo- domorphes comprennent les Sauropodes (Diplodocus, Brachiosaurus...), fossiles célébrissimes du fait de leur statut de plus grands vertébrés terrestres, avec toutes les conséquences paléobiologiques intéressantes posées par ce gigantisme. On le sait (voir Annuaire 2000-2001 pp. 433-437), Théropodes (incl. oiseaux) et Sauropodomorphes constituent ensemble une unité naturelle ou clade (Sau- rischia ou Sauripelvia) au sein des Dinosauria, ceux-ci se situant parmi les Ornithosuchia au sein des Archosauria. L’autre grand rameau évolutif des Archo- saures, les Pseudosuchia, comprend les Crocodylia et se poursuit donc, avec ces derniers, dans la nature actuelle. Nous avons déjà envisagé (ibid. pp. 437-439) l’évolution initiale des Pseudosuchia au Trias, il nous restait donc à poursuivre l’analyse de l’évolution de cette ligné depuis la base du Jurassique jusqu’à l’actuel, ce par quoi j’ai commencé les leçons de cette année. Comme d’habitude, j’ai tenté, dans cette série d’exposés, de faire distinguer clairement les connaissances acquises des interprétations et inférences plus ou moins directes concernant la phylogénie comme la paléobiologie. Au-delà des données anatomiques factuelles présentées les exposés ont donc systématique- ment croisé trois composantes, dans une perspective critique : a/ d’une part, les 462 ARMAND DE RICQLÈS données paléogéographiques, stratigraphiques et paléoclimatologiques concernant le Mésozoïque et ses gisements à vertébrés continentaux, b/ d’autre part, les données d’histoire de la paléontologie et de la systématique portant sur la décou- verte, la compréhension, la classification et la nomenclature « traditionnelle » des groupes envisagés, c/ enfin, le renouveau des conceptions, accompli au cours des quinze dernières années grâce à l’application de la méthode cladistique, abou- tissant à des hypothèses phylogénétiques argumentées, sinon toujours robustes, renouveau désormais associéàcelui des principes et de la pratique de la nomen- clature systématique (définition phylogénétique des taxons). Pseudosuchia : Crocodyliforma : Mesoeucrocodylia. La suite de l’analyse de l’évolution de ce clade a occupé les cinq premières leçons. A` l’inverse du déroulement général des exposés selon le décours temporel « normal », je suis parti cette fois de la considération « herpétologique » de la faune actuelle des crocodiliens (biologie, éco-ethologie, répartition, systématique...) pour « remonter le temps » en direction des abondants documents fossiles relatant l’évolution des crocodiles « modernes » (Eusuchia) à partir du Crétacé supérieur et au cours du Cénozoïque. Cette approche a permis de confronter les données paléontologiques et néontologiques (particulièrement moléculaires) pour la reconstruction phylogé- nétique. A` la suite de quoi, j’ai repris le décours temporel « normal » pour traiter l’évolution des crocodiliens plus anciens (principalement Mesoeucrocodylia très divers d’âges Jurassique et Crétacé) succédant aux formes triasiques (Sphenosu- chia, etc.) déjàévoquées il y a deux ans. Dans la faune actuelle, les crocodiliens ne sont représentés que par une faible biodiversité : crocodiles, caïmans et gavials comprennent 22 espèces réparties dans 8 genres et 3 Familles. Écologiquement, ils constituent toutefois un groupe de superprédateurs très bien adaptés dans leurs environnements et seule la pres- sion humaine limite actuellement leur succès dans tous les milieux intertropicaux riches en eaux douces. J’ai d’abord passé en revue les principales caractéristiques anatomo-physiologiques du groupe (appareil circulatoire et respiratoire, système nerveux central, appareil locomoteur et locomotion...) dont certains aspects constituent des apomorphies soit au niveau des crocodiliens eux-mêmes, soit peut être plus généralement au niveau des Archosaures. La répartition géographique est liée à des caractéristiques spécifiques (adaptation au froid, tolérance à la salinité...). Morphologies crâniennes (brévi-, méso- ou longirostrie...) et dentaires sont manifestement liées à la fois à l’écologie (plus ou moins terrestre) et au régime alimentaire (plus ou moins piscivore). La reproduction, caractérisée par le déterminisme environnemental du sexe, fait intervenir des moeurs assez variées au niveau spécifique, avec construction de divers types de nids et des interactions importantes entre générations. La croissance montre une plasticité considérable, à la fois au plan génétique et en fonction des conditions de l’environnement. Compte tenu des défis actuels posés par la conservation de la biodiversité j’ai détaillé la situation, globalement très alarmante chez les crocodiliens, avec toute- BIOLOGIE HISTORIQUE ET ÉVOLUTIONNISME 463 fois quelques exemples assez encourageants de restauration (Gavial) ou d’exploi- tations raisonnées des populations. La systématique traditionnelle en Alligatoridés (4 genres, 7 espèces), Crocody- lidés (3 genres, 14 espèces) et Gavialidés (1 genre, une espèce), progressivement établie au XIXe siècle, a été confortée par les études morphologiques récentes mais pose toutefois des problèmes, compte tenu de l’enracinement ancien des taxons actuels (Crétacé) et de la complexité de leur histoire phylogénétique et biogéographique au cours du Cénozoïque. De fait, l’introduction depuis 15 ans des données moléculaires permet la constitution de nouveaux « jeux de données » indépendants de la morphologie. La confrontation, pour les mêmes taxons, de ces jeux de données indépendants est évidemment du plus haut intérêt aux plans théorique et méthodologique. J’ai largement exposé, à cet égard, la magistrale synthèse de Chr. Brochu (1997). On peut en retenir ici la remarquable similitude des résultats phylogénétiques issus de l’analyse indépendante des jeux de données morphologiques et moléculaires, sauf sur un seul point : la position du gavial. Le désaccord porte sur deux aspects : a/ un problème de topologie d’arbres : où se place le gavial (et ses parents éteints) sur le cladogramme des Crocodylia ? ; b/ un problème temporel : quand la lignée du gavial a-t-elle divergée de ses plus proches parents actuels ? Pour les morphologistes (et paléontologues) le gavial actuel représente une lignée indépendante détachéedèsleCrétacé supérieur (70 MA), et les Crocody- lidés et Alligatoridés sont plus proches parents les uns des autres que chacun d’eux ne l’est des Gavialidés. Pour les molécularistes, au contraire, le gavial forme un clade avec le faux gavial (Tomistoma), clade plus apparenté aux Croco- dylidésqu’aux Alligatoridés. De plus Gavialis et Tomistoma n’auraient divergé qu’au Pliocène, au plus tôt au Miocène (15 MA). Du fait de la qualité et de la quantité des jeux de données morphologiques (y compris paléontologiques) et moléculaires, par ailleurs d’accord sur l’ensemble du cladogramme, la « non congruence » (contradiction) entre résultats issus de ces jeux de données différents à propos du gavial constitue un véritable cas d’école, illustrant bon nombre de problèmes et biais méthodologiques en recons- truction phylogénétique, que l’on a détaillé au cours des exposés. Dans le cas présent, la combinaison de l’ensemble des données en une seule matrice (Poe 1996) semble discutable. En effet, la combinaison de jeux de données contradic- toires mais chacun solidement soutenu est incapable de produire une phylogénie fiable. En l’occurrence, la permanence et la force avec lesquelles les divers jeux de données sont en contradiction quant à la place de Gavialis et son temps de divergence indiquent un conflit lié aux données elles-mêmes. A` l’heure actuelle, le problème ne peut être considéré comme résolu. Les Pseudosuchia post-Triasiques appartiennent tous au groupe apical des Cro- codyliformes. La systématique évolutionniste y distinguait traditionnellement trois « grades » ou « paliers évolutifs » de plus en plus « progressifs » : Protosu- 464 ARMAND DE RICQLÈS chia, Mesosuchia et Eusuchia se succédant du Trias supérieur à l’actuel, point de vue réfuté par l’analyse cladistique, du fait de la paraphylie intrinsèque au concept de Mesosuchia. Comme on l’a vu (ibid. p. 438), les Protosuchia semblent bien constituer un petit groupe naturel ou clade, connu du Trias supérieur et du Jurassique inférieur. Ils s’opposent à l’ensemble plus dérivé des « Mesoeucroco- dylia », incluant tous les autres Crocodyliformes dont les formes actuelles. L’ana- lyse phylogénétique des Mesoeucrocodylia est très complexe et un consensus général n’est pas encore établi. On se bornera ici à un canevas simplifié (Fig. 1), sans entrer dans la discussion des nombreuses hypothèses alternatives qui ont été présentées au cours des exposés. Figure 1 : Cladogramme « consensus » (simplifié) des Crocodyliformes. Les taxons monophylétiques 1 — 5 (ouvertures de parenthèses) sont définis phylogénétiquement comme branches (« stem defined »), voir texte : 1 = Mesoeucrocodylia, 2 = Thalattosu- chia, 3 = Metasuchia, 4 = Neosuchia, 5 = Eusuchia, (*) = taxons longirostres
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